, Volume 38, Issue 6, pp 1133–1145 | Cite as

The Effects of Area and Habitat Heterogeneity on Bird Richness and Composition in High Elevation Wetlands (“Bofedales”) of the Central Andes of Peru

  • Grace P. Servat
  • Renzo Alcocer
  • Melvi V. Larico
  • Magaly E. Olarte
  • Reynaldo Linares-Palomino
  • Alfonso Alonso
  • Karim Ledesma
Wetlands in the Developing World


We assessed area and habitat heterogeneity effects on avian richness and composition in bofedales that differed in size and microhabitat diversity. We analyzed data collected in 2 seasons and 24 bofedales using General Linear Models, Ordinary Least Square models to establish the relationship of predictor variables on richness and Akaike Information Criterion for model selection. We evaluate composition classifying species into groups using Bray Curtis ordination, followed by Multiple Response Permutation Procedure to test for differences among groups, and Indicator Species Analysis to identify species. Bofedales differed in richness (F = 5.1, p < 0.001) and microhabitat diversity (F = 23.4, p < 0.001), but no seasonal differences emerged (p > 0.05). The best model indicates that 54% of variance in richness was explained by area and microhabitat diversity, however, a tendency to decrease in microhabitat diversity as area increases, suggests a relatively more important role of area. Results are supported by composition, as microhabitats not only differed pairwise (T = −94.14, A = 0.601, p < 0.001) and had significant indicator species (p < 0.05), but because its differential contribution to richness, as some microhabitats were more speciose than others. As such, few species-rich microhabitats may contribute more to richness than many species-poor ones which is not predicted by the habitat heterogeneity hypothesis. Disentangling the influence of area and habitat heterogeneity on species richness is important to establish conservation priorities that ensure bofedales integrity under imminent climate change.


Diversity Microhabitats Cover Cushion bogs Indicator species 


Evaluamos el efecto del área y la heterogeneidad del hábitat en la riqueza y composición de aves en bofedales que difieren en tamaño y diversidad de microhábitats. Los datos recopilados en 2 estaciones y 24 bofedales fueron analizados usando Modelos Generales Lineales, Modelos de Mínimos Cuadrados Ordinarios para establecer la relación entre las variables predictivas y la riqueza, y el Criterio de Información de Akaike para seleccionar los modelos. Evaluamos la composición de especies clasificándolas en grupos con el Ordenamiento de Bray Curtis, seguido por el Análisis de Permutación de Respuesta Múltiple para detectar diferencias entre los grupos, y el Análisis de Especies Indicadoras para identificar las especies. Los bofedales difieren en riqueza (F = 5.1, p < 0.001) y diversidad de microhábitats (F = 23.4, p < 0.001), pero no hallamos diferencias estacionales (p > 0.05). El modelo seleccionado indica que el área y la diversidad de microhábitats explican 54% de la varianza en la riqueza, sin embargo, encontramos una tendencia inversa entre la diversidad de microhábitats y el área, la cual sugiere un papel relativamente más importante del área en la riqueza de especies. Nuestros resultados son respaldados por los datos de composición, ya que los microhábitats no sólo fueron diferentes en comparaciones pareadas (T = −94.14, A = 0.601, p < 0.001) y estuvieron representados significativamente por especies indicadoras (p < 0.05), sino que contribuyeron diferencialmente con la riqueza. Así, pocos microhábitats ricos en especies contribuirían más a la riqueza que varios microhábitats pobres en especies lo cual no concuerda con las predicciones de la hipótesis de heterogeneidad del hábitat. Determinar la influencia que el área y la heterogeneidad tienen en la riqueza de especies es importante para establecer prioridades de conservación que garanticen la integridad de los bofedales ante el inminente cambio climático.

Palabras clave

Diversidad Microhábitats Cobertura  Bofedales Especies indicadoras 



This study is a contribution of the Biodiversity Monitoring and Assessment Program (BMAP) from the Center of Conservation and Sustainability of the Smithsonian Conservation Biology Institute and PERU-LNG. We are thankful to all BMAP personnel in Lima, Ayacucho, and Washington DC, for their support with field logistics throughout the study, and T. Erwin for reviewing an early version of the manuscript. We specially thank past and present administrators of the program for their contribution at different stages of the project. Authorizations to conduct fieldwork were issued by the Direccion General de Flora y Fauna Silvestre (RD 405–2012-AG-DGFFS-GDEFFS). This is contribution No. 43 of the Peru Biodiversity Program of the Smithsonian Conservation Biology Institute.


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Copyright information

© Society of Wetland Scientists 2017

Authors and Affiliations

  • Grace P. Servat
    • 1
    • 2
  • Renzo Alcocer
    • 3
  • Melvi V. Larico
    • 3
  • Magaly E. Olarte
    • 4
  • Reynaldo Linares-Palomino
    • 5
  • Alfonso Alonso
    • 1
  • Karim Ledesma
    • 5
  1. 1.Center for Conservation and SustainabilitySmithsonian Conservation Biology InstituteWashingtonUSA
  2. 2.Departamento de OrnitologíaMuseo de Historia Natural de la Universidad Nacional Mayor de San MarcosLimaPeru
  3. 3.Museo de Historia Natural de la Universidad Nacional de San Agustín de ArequipaArequipaPeru
  4. 4.Museo de Historia Natural de la Universidad Nacional San Antonio Abad del Cusco, Paraninfo Universitario, Plaza de ArmasCuscoPeru
  5. 5.Center for Conservation and SustainabilitySmithsonian Conservation Biology InstituteLimaPeru

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